Linking fine root morphology, hydraulic functioning and shade tolerance of trees.

Background and Aims: Understanding root traits and their trade-off with other plant processes is important for understanding plant functioning in natural ecosystems as well as agricultural systems. The aim of the present study was to determine the relationship between root morphology and the hydraulic characteristics of several orders of fine roots (<2 mm) for species differing in shade tolerance (low, moderate and high). Methods: The morphological, anatomical and hydraulic traits across five distal root orders were measured in species with different levels of shade tolerance and life history strategies. The species studied were Acer negundo, Acer rubrum, Acer saccharum, Betula alleghaniensis, Betula lenta, Quercus alba, Quercus rubra, Pinus strobus and Pinus virginiana. Key Results: Compared with shade-tolerant species, shade-intolerant species produced thinner absorptive roots with smaller xylem lumen diameters and underwent secondary development less frequently, suggesting that they had shorter life spans. Shade-tolerant species had greater root specific hydraulic conductance among these roots due to having larger diameter xylems, although these roots had a lower calculated critical tension for conduit collapse. In addition, shade-intolerant species exhibited greater variation in hydraulic conductance across different root growth rings in woody transport roots of the same root order as compared with shade-tolerant species. Conclusions: Plant growth strategies were extended to include root hydraulic properties. It was found that shade intolerance in trees was associated with conservative root hydraulics but greater plasticity in number of xylem conduits and hydraulic conductance. Root traits of shade-intolerant species were consistent with the ability to proliferate roots quickly for rapid water uptake needed to support rapid shoot growth, while minimizing risk in uncertain environments.

Light energy partitioning, photosynthetic efficiency and biomass allocation in invasive Prunus serotina and native Quercus petraea in relation to light environment, competition and allelopathy.

This study addressed whether competition under different light environments was reflected by changes in leaf absorbed light energy partitioning, photosynthetic efficiency, relative growth rate and biomass allocation in invasive and native competitors. Additionally, a potential allelopathic effect of mulching with invasive Prunus serotina leaves on native Quercus petraea growth and photosynthesis was tested. The effect of light environment on leaf absorbed light energy partitioning and photosynthetic characteristics was more pronounced than the effects of interspecific competition and allelopathy. The quantum yield of PSII of invasive P. serotina increased in the presence of a competitor, indicating a higher plasticity in energy partitioning for the invasive over the native Q. petraea, giving it a competitive advantage. The most striking difference between the two study species was the higher crown-level net CO2 assimilation rates (Acrown) of P. serotina compared with Q. petraea. At the juvenile life stage, higher relative growth rate and higher biomass allocation to foliage allowed P. serotina to absorb and use light energy for photosynthesis more efficiently than Q. petraea. Species-specific strategies of growth, biomass allocation, light energy partitioning and photosynthetic efficiency varied with the light environment and gave an advantage to the invader over its native competitor in competition for light. However, higher biomass allocation to roots in Q. petraea allows for greater belowground competition for water and nutrients as compared to P. serotina. This niche differentiation may compensate for the lower aboveground competitiveness of the native species and explain its ability to co-occur with the invasive competitor in natural forest settings.

Drivers of radial growth and carbon isotope discrimination of bur oak (Quercus macrocarpa†Michx.) across continental gradients in precipitation, vapour pressure deficit and irradiance.

Tree-ring characteristics are commonly used to reconstruct climate variables, but divergence from the assumption of a single biophysical control may reduce the accuracy of these reconstructions. Here, we present data from bur oaks (Quercus macrocarpa Michx.) sampled within and beyond the current species bioclimatic envelope to identify the primary environmental controls on ring-width indices (RWIs) and carbon stable isotope discrimination (Δ(13) C) in tree-ring cellulose. Variation in Δ(13) C and RWI was more strongly related to leaf-to-air vapour pressure deficit (VPD) at the centre and western edge of the range compared with the northern and wettest regions. Among regions, Δ(13) C of tree-ring cellulose was closely predicted by VPD and light responses of canopy-level Δ(13) C estimated using a model driven by eddy flux and meteorological measurements (R(2) = 0.96, P = 0.003). RWI and Δ(13) C were positively correlated in the drier regions, while they were negatively correlated in the wettest region. The strength and direction of the correlations scaled with regional VPD or the ratio of precipitation to evapotranspiration. Therefore, the correlation strength between RWI and Δ(13) C may be used to infer past wetness or aridity from paleo wood by determining the degree to which carbon gain and growth have been more limited by moisture or light.

Relationship between photochemical reflectance index and leaf ecophysiological and biochemical parameters under two different water statuses: towards a rapid and efficient correction method using real-time measurements.

The use of the photochemical reflectance index (PRI) as a promising proxy of light use efficiency (LUE) has been extensively studied, and some issues have been identified, notably the sensitivity of PRI to leaf pigment composition and the variability in PRI response to LUE because of stress. In this study, we introduce a method that enables us to track the short-term PRI response to LUE changes because of photosynthetically active radiation (PAR) changes. The analysis of these short-term relationships between PRI and LUE throughout the growing season in two species (Quercus robur L. and Fagus sylvatica L.) under two different soil water statuses showed a clear change in PRI response to LUE, which is related to leaf pigment content. The use of an estimated or approximated PRI0, defined as the PRI of perfectly dark-adapted leaves, allowed us to separate the PRI variability due to leaf pigment content changes and the physiologically related PRI variability over both daily (PAR-related) and seasonal (soil water content-related) scales. The corrected PRI obtained by subtracting PRI0 from the PRI measurements showed a good correlation with the LUE over both of the species, soil water statuses and over the entire growing season.

Three pools of zeaxanthin in Quercus coccifera leaves during light transitions with different roles in rapidly reversible photoprotective energy dissipation and photoprotection.

Under excess light, the efficient PSII light-harvesting antenna is switched into a photoprotected state in which potentially harmful absorbed energy is thermally dissipated. Changes occur rapidly and reversibly, enhanced by de-epoxidation of violaxanthin (V) to zeaxanthin (Z). This process is usually measured as non-photochemical quenching (NPQ) of chlorophyll (Chl) fluorescence. Using instrumentation for instantaneous leaf freezing, NPQ, spectral reflectance, and interconversions within the xanthophyll cycle with time resolution of seconds were recorded from Quercus coccifera leaves during low light (LL) to high light (HL) transitions, followed by relaxation at LL. During the first 30 s of both the LL to HL and HL to LL transitions, no activity of the xanthophyll cycle was detected, whereas 70-75% of the NPQ was formed and relaxed, respectively, by that time, the latter being traits of a rapidly reversible photoprotective energy dissipation. Three different Z pools were identified, which play different roles in energy dissipation and photoprotection. In conclusion, ΔpH was crucial to NPQ formation and relaxation in Q. coccifera during light transitions. Only a minor fraction of Z was associated to quenching, whereas the largest Z pool was not related to thermal dissipation. The latter is proposed to participate in photoprotection acting as antioxidant.

Light and competition gradients fail to explain the coexistence of shade-tolerant Fagus sylvatica and shade-intermediate Quercus petraea seedlings.

BACKGROUND AND AIMS: The coexistence of forest tree species has often been linked to differences among species in terms of their response to light availability during the regeneration stage. From this perspective, species coexistence results from growth-growth or mortality-growth trade-offs along spatial light gradients. Experimental evidence of growth-growth trade-offs in natural conditions is sparse due to various confounding factors that potentially hinder the relationship. This study examined growth hierarchies along light gradients between two tree species with contrasting shade tolerance by controlling potential confounding factors such as seedling size, seedling status, seedling density and species composition. METHODS: Natural regenerated shade-tolerant Fagus sylvatica and shade-intermediate Quercus petraea seedlings were used, and growth rankings over a 4-year period were compared in 8- to 10-year-old tree seedlings. KEY RESULTS: No rank reversal occurs between the two species along the light gradient, or along the density, mixture or seedling size gradients. The shade-tolerant species was always the more competitive of the two. Pronounced effects of initial size on seedling growth were observed, whereas the effects of light and competition by neighbours were of secondary importance. The paramount effect of size, which results from the asymmetric nature of interseedling competition, gives a strong advantage to tall seedlings over the long term. CONCLUSIONS: This study extends previous efforts to identify potential drivers of rank reversals in young tree mixtures. It does not support the classical assumption that spatial heterogeneity in canopy opening explains the coexistence of the two species studied. It suggests that spatial variation in local size hierarchies among seedlings that may be caused by seedling emergence time or seedling initial performance is the main driver of the dynamics of these mixed stands.

Hydraulic conductivity of red oak (Quercus rubra L.) leaf tissue does not respond to light.

The permeability of leaf tissue to water has been reported to increase under illumination, a response reputed to involve aquaporins. We studied this 'light response' in red oak (Quercus rubra L.), the species in which the phenomenon was first detected during measurements of leaf hydraulic conductance with the high-pressure flow meter (HPFM). In our HPFM measurements, we found that pre-conditioning leaves in darkness was not sufficient to bring them to their minimum conductance, which was attained only after an hour of submersion and pressurization. However, pre-conditioning leaves under anoxic conditions resulted in an immediate reduction in conductance. Leaves light- and dark-acclimated while on the tree showed no differences in the time course of HPFM measurement under illumination. We also studied the effect of light level and anoxia on rehydration kinetics, finding that anoxia slowed rehydration, but light had no effect either in the lab (rehydration under low light, high humidity) or on the tree (acclimation under high light, 10 min of dark prior to rehydration). We conclude that the declines in conductance observed in the HPFM must involve a resistance downstream of the extracellular air space, and that in red oak the hydraulic conductivity of leaf tissue is insensitive to light.

Temporal variation in leaf nitrogen partitioning of a broad-leaved evergreen tree, Quercus myrsinaefolia.

We examined temporal changes in the amount of nitrogenous compounds in leaves from the outer and inner parts of the crown of Quercus myrsinaefolia growing in a seasonal climate. Throughout the leaf life span, metabolic protein and Rubisco content closely correlated with total nitrogen content, while structural protein content was relatively stable after full leaf expansion. Chlorophyll content was affected by shading as well as total nitrogen content in outer leaves that were overtopped by new shoots in the second year. Outer leaves showed a large seasonal variation in photosynthetic nitrogen-use efficiency (PNUE; the light-saturated photosynthetic rate per unit leaf nitrogen content) during the first year of their life, with PNUE decreasing from the peak in summer towards winter. Outer and inner leaves both showed age-related decline in PNUE in the second year. There were no such drastic changes in leaf nitrogen partitioning that could explain seasonal and yearly variations in PNUE. Nitrogen resorption occurred in overwintering leaves in spring. Metabolic protein explained the majority of nitrogen being resorbed, whereas structural protein, which was low in degradability, contributed little to nitrogen resorption.

Effects of light spectra and 15N pulses on growth, leaf morphology, physiology, and internal nitrogen cycling in Quercus variabilis Blume seedlings.

Light spectra of sunlight transmittance can generate an interactive effect with deposited nitrogen (N) on regenerated plants across varied shading conditions. Total N content in understory plants can be accounted for by both exogeneous and endogenous sources of derived N, but knowledge about the response of inner N cycling to interactive light and N input effects is unclear. We conducted a bioassay on Chinese cork oak (Quercus variabilis Blume) seedlings subjected to five-month N pulsing with 15NH4Cl (10.39 atom %) at 120 mg 15N plant-1 under the blue (48.5% blue, 33.7% green, and 17.8% red), red (14.6% blue, 71.7% red, 13.7% green), and green (17.4% blue, 26.2% red, 56.4% green) lighting-spectra. Half of the seedlings were fed twice a week using a 250 ppm N solution with micro-nutrients, while the other half just received distilled water. Two factors showed no interaction and neither affected growth and morphology. Compared to the red-light spectrum, that in blue light increased chlorophyll and soluble protein contents and glutamine synthetase (GS) activity, root N concentration, and N derived from the pulses. The green-light spectrum induced more biomass allocation to roots and a higher percentage of N derived from internal reserves compared to the red-light spectrum. The 15N pulses reduced the reliance on N remobilization from acorns but strengthened shoot biomass, chlorophyll content, GS activity, and N concentration. In conclusion, light spectrum imposed an independent force from external N pulse to modify the proportion of N derived from internal sources in total N content in juvenile Q. variabilis.

Aquaporin gene expression and apoplastic water flow in bur oak (Quercus macrocarpa) leaves in relation to the light response of leaf hydraulic conductance.

It has previously been shown that hydraulic conductance in bur oak leaves (Quercus macrocarpa Michx.), measured with the high pressure flow meter technique (HPFM), can significantly increase within 30 min following exposure to high irradiance. The present study investigated whether this increase could be explained by an increase in the cell-to-cell pathway and whether the response is linked to changes in the transcript level corresponding to aquaporin genes. Four cDNA sequences showing high similarity to members of the aquaporin gene family from other plant species were characterized from bur oak leaves and the expression levels of these cDNA sequences were examined in leaves by quantitative real-time PCR (QRT-PCR). No change was found in the relative transcript abundance corresponding to these four putative aquaporin genes in leaves with light-induced high hydraulic conductance (exposed to high irradiance) compared to leaves with low hydraulic conductance (exposed to low irradiance). However, in sun leaves that were exposed to different light levels prior to leaf collection (full sunlight, shade, and covered with aluminium foil for 16 h), the relative transcript levels of two of the putative aquaporin genes increased several-fold in shaded leaves compared to the sun-exposed or covered leaves. When the leaves were pressure-infiltrated with the apoplastic tracer dye trisodium 3-hydroxy-5,8,10-pyrenetrisulphonate (PTS(3), 0.02%), there was no change in the PTS(3) concentration of leaf exudates collected in ambient light or in high irradiance, but there was a small apoplastic acidification. There was also no change in PTS(3) concentration between the leaves infiltrated under high irradiance with 0.02% PTS(3) or with 0.1 mM HgCl(2) in 0.02% PTS(3). The results suggest that the putative aquaporin genes that were identified in the present study probably do not play a role in the light responses of hydraulic conductance at the transcript level, but they may function in regulating water homeostasis in leaves adapted to different light conditions. In addition, it is shown that high irradiance induced changes in the pH of the apoplast and that there does not appear to be a significant shift to the cell-to-cell mediated water transport in bur oak leaves exposed to high irradiance as measured by the apoplastic tracer dye.

Developmental changes in spatial distribution of in vivo fluorescence and epidermal UV absorbance over Quercus petraea leaves.

BACKGROUND AND AIMS: Epidermal phenolic compounds (mainly flavonoids) constitute a vital screen that protects the leaf from damage by natural ultraviolet (UV) radiation. The effectiveness of epidermal UV-screening depends on leaf anatomy, the content of UV-screening compounds and their spatial uniformity over the leaf area. To investigate in vivo the spatial pattern of the epidermal UV-screen during leaf development, a fluorescence imaging method was developed to map the epidermal UV-absorbance at a microscopic scale. This study was done on oak (Quercus petraea) leaves that were used as a model of woody dicotyledonous leaves. METHODS: The leaf development of 2-year-old trees, grown outdoors, was monitored, at a macroscopic scale, by in vivo measurements of chlorophyll content per unit area and epidermal UV-absorbance using two optical leaf-clip meters. The distribution of pigments within leaves was assessed in vivo spectroscopically. The microscopic images of UV-induced fluorescence and UV-absorbance acquired in vivo during leaf development were interpreted from spectral characteristics of leaves. KEY RESULTS: At a macroscopic scale, epidermal UV-absorbance was high on the upper leaf side during leaf development, while it increased on the lower leaf side during leaf expansion and reached the adaxial value at maturity. At a microscopic scale, in immature leaves, for both leaf sides, the spatial distribution of epidermal UV-absorbance was heterogeneous, with a pattern depending on the flavonoid content of vacuoles in developing epidermal cells. At maturity, epidermal UV-absorbance was uniform. CONCLUSIONS: The spatial pattern of epidermal UV-screen over the area of oak leaves is related to leaf anatomy during development. In vivo spectroscopy and fluorescence imaging of the leaf surface showed the distribution of pigments within the leaf and hence can provide a tool to monitor optically the leaf development in nature.

Leaf biochemical adjustments in two Mediterranean resprouter species facing enhanced UV levels and reduced water availability before and after aerial biomass removal.

Effects of supplemented UV radiation and diminished water supply on the leaf concentrations of phenols and antioxidants of two Mediterranean resprouter species, Arbutus unedo and Quercus suber, were assessed before and after entire aerial biomass removal. Potted seedlings of both species were grown outdoors for 8 months with enhanced UV-A + UV-B, enhanced UV-A or ambient UV, in combination with two watering conditions (field capacity or watering reduction). After this period, all aerial biomass was removed and new shoots (resprouts) developed for a further 8 months under the two treatments. In general, the investment in leaf phenols was substantially greater in A. unedo than in Q. suber, while Q. suber allocated more resources to non-phenolic antioxidants (ascorbate and glutathione). In response to enhanced UV-B radiation, Q. suber leaves rose their UV-screening capacity mainly via accumulation of kaempferols, accompanied by an increased concentration of rutins, being these effects exacerbated under low-watering conditions. Conversely, A. unedo leaves responded to UV-B radiation reinforcing the antioxidant machinery by increasing the overall amount of flavonols (especially quercetins) in seedlings, and of ascorbate and glutathione, along with catalase activity, in resprouts. Nevertheless, UV effects on the amount/activity of non-phenolic antioxidants of A. unedo resprouts were modulated by water supply. Indeed, the highest concentration of glutathione was found under the combination of enhanced UV-B radiation and reduced watering, suggesting an enlargement of the antioxidant response in A. unedo resprouts. Different biochemical responses to enhanced UV and drier conditions in seedlings and resprouts of these two species might modulate their competitive interactions in the near future.

Methanol emissions from deciduous tree species: dependence on temperature and light intensity.

Methanol emissions from several deciduous tree species with predominantly mature leaves were measured under laboratory and field conditions. The emissions were modulated by temperature and light. Under constant light conditions in the laboratory, methanol emissions increased with leaf temperature, by up to 12% per degree. At constant temperatures, emissions doubled when light intensity (PAR) increased from darkness to 800 micromol x m(-2) x s(-1). A phenomenological description of light and temperature dependencies was derived from the laboratory measurements. This description was successfully applied to reproduce the diel cycle of methanol emissions from an English oak measured in the field. Labelling experiments with (13)CO(2) provided evidence that less than 10% of the emitted methanol was produced de novo by photosynthesis directly prior to emission. Hence, the light dependence of the emissions cannot be explained by instantaneous production from CO(2) fixation. Additional experiments with selective cooling of plant roots indicated that a substantial fraction of the emitted methanol may be produced in the roots or stem and transported to stomata by the transpiration stream. However, the transpiration stream cannot be considered as the main factor that determines methanol emissions by the investigated plants.

Leaf physiological versus morphological acclimation to high-light exposure at different stages of foliar development in oak.

We investigated light acclimation in seedlings of the temperate oak Quercus petraea (Matt.) Liebl. and the co-occurring sub-Mediterranean oak Quercus pyrenaica Willd. Seedlings were raised in a greenhouse for 1 year in either 70 (HL) or 5.3% (LL) of ambient irradiance of full sunlight, and, in the following year, subsets of the LL-grown seedlings were transferred to HL either before leaf flushing (LL-HLBF plants) or after full leaf expansion (LL-HLAF plants). Gas exchange, chlorophyll a fluorescence, nitrogen fractions in photosynthetic components and leaf anatomy were examined in leaves of all seedlings 5 months after plants were moved from LL to HL. Differences between species in the acclimation of LL-grown plants to HL were minor. For LL-grown plants in HL, area-based photosynthetic capacity, maximum rate of carboxylation, maximum rate of electron transport and the effective photochemical quantum yield of photosystem II were comparable to those for plants grown solely in HL. A rapid change in nitrogen distribution among photosynthetic components was observed in LL-HLAF plants, which had the highest photosynthetic nitrogen-use efficiency. Increases in mesophyll thickness and dry mass per unit area governed leaf acclimation in LL-HLBF plants, which tended to have less nitrogen in photosynthetic components and a lower assimilation potential per unit of leaf mass or nitrogen than LL-HLAF plants. The data indicate that the phenological state of seedlings modified the acclimatory response of leaf attributes to increased irradiance. Morphological adaptation of leaves of LL-HLBF plants enhanced photosynthetic capacity per unit leaf area, but not per unit leaf dry mass, whereas substantial redistribution of nitrogen among photosynthetic components in leaves of LL-HLAF plants enhanced both mass- and area-based photosynthetic capacity.

Water use and carbon exchange of red oak- and eastern hemlock-dominated forests in the northeastern USA: implications for ecosystem-level effects of hemlock woolly adelgid.

Water use and carbon exchange of a red oak-dominated (Quercus rubra L.) forest and an eastern hemlock-dominated (Tsuga canadensis L.) forest, each located within the Harvard Forest in north-central Massachusetts, were measured for 2 years by the eddy flux method. Water use by the red oak forest reached 4 mm day(-1), compared to a maximum of 2 mm day(-1) by the eastern hemlock forest. Maximal carbon (C) uptake rate was also higher in the red oak forest than in the eastern hemlock forest (about 25 versus 15 micromol m(-2) s(-1)). Sap flux measurements indicated that transpiration of red oak, and also of black birch (Betula lenta L.), which frequently replaces eastern hemlock killed by hemlock woolly adelgid (Adelges tsugae Annand.), were almost twice that of eastern hemlock. Despite the difference between species in maximum summertime C assimilation rate, annual C storage of the eastern hemlock forest almost equaled that of the red oak forest because of net C uptake by eastern hemlock during unusually warm fall and spring weather, and a near-zero C balance during the winter. Thus, the effect on C storage of replacing eastern hemlock forest with a forest dominated by deciduous species is unclear. Carbon storage by eastern hemlock forests during fall, winter and spring is likely to increase in the event of climate warming, although this may be offset by C loss during hotter summers. Our results indicate that, although forest water use will decrease immediately following eastern hemlock mortality due to the hemlock woolly adelgid, the replacement of eastern hemlock by deciduous species such as red oak will likely increase summertime water use over current rates in areas where hemlock is a major forest species.

Light response of hydraulic conductance in bur oak (Quercus macrocarpa) leaves.

A four- to seven-fold enhancement of leaf hydraulic conductance by light has been reported in three temperate tree species. The enhancement occurs in the liquid-flow pathway between the petiole and the site of water evaporation. The enhancement occurs within 1 h, and dissipates in darkness over a period of 1 to 10 h depending on species. Here we report light-induced enhancement of leaf hydraulic conductance in a fourth species, bur oak (Quercus macrocarpa Michx.), the dependence of the effect on light flux and color, its absence in leaves of seedlings, and the impact on the response of leaf vein severance and several metabolic inhibitors. The light response of leaf hydraulic conductance approached saturation at a photosynthetic photon flux of 150 mumol m(-2) s(-1). Hydraulic enhancement was greater in response to blue and green light than to visible radiation of longer wavelengths, although at the same irradiance, the response to white light was greater than to light of any single color. Atrazine (a photosystem II inhibitor), fusicoccin (which stimulates plasma membrane-bound H(+)-ATPase) and HgCl(2) (an aquaporin blocker) reduced the light response of leaf lamina hydraulic conductance. When 2-mercaptoethanol was added following mercury treatment, the light response was totally suppressed. Our results are consistent with the notion that the effect of light on leaf lamina hydraulic conductance is controlled by factors acting outside the leaf veins, possibly through light-induced changes in membrane permeability of either mesophyll or bundle sheath cells, or both.

Nitrogen deposition does not affect the impact of shade on Quercus acutissima seedlings.

Light and atmospheric nitrogen (N) deposition are among the important environmental factors influencing plant growth and forest regeneration. We used Quercus acutissima, a dominant broadleaf tree species native to the deciduous forests of Northern China, to study the combined effects of light exposure and N addition on leaf physiology and individual plant growth. In the greenhouse, we exposed Quercus acutissima seedlings to one of two light conditions (8% and 80% of full irradiation) and one of three N treatments (0, 6, and 12 g N m-2 y-1). After 87 d, we observed that nitrogen deposition had no significant effects on the seedlings regardless of light exposure. In addition, shade significantly reduced plant height, basal diameter, leaf number, total biomass, gas exchange capacity, and carbohydrate content. In contrast, however, shade significantly increased the amount of photosynthetic pigment, above-ground biomass allocation, and specific leaf area. There was also a hierarchical plasticity among the different seedling characteristics. Compared to traits of growth, biomass, biomass allocation and leaf morphology, the leaf physiology, including photosynthetic pigment, gas exchange, carbohydrate, and PUNE, is more sensitive to light conditions. Among the biomass allocation parameters, the leaf and root mass ratios had a relatively low phenotypic plasticity. The seedlings had high foliar physiological plasticity under various light conditions. Nevertheless, we recommend high irradiance to maintain vigorous seedling growth and, in turn, promote the restoration and reconstruction of vegetation.

Response of transglutaminase activity and bound putrescine to changes in light intensity under natural or controlled conditions in Quercus ilex leaves.

In order to further study a previously observed relationship between polyamine (PA) content and changes in irradiation, we examined the level of free and bound PAs, the activity of transglutaminase (TGase, EC 2.3.2.13) and chlorophyll fluorescence in holm oak (Quercus ilex L.) leaves in response to different levels of light intensity and amount. A diurnal trend of free and bound putrescine (F-Put and B-Put, respectively) and TGase activity was observed in plants under natural conditions in the forest, with the highest value corresponding to the maximum light intensity and amount of light received by the leaves. In another set of experiments, potted Q. ilex plants in experimental fields were subjected to a range of periods of natural photosynthetic photon flux density (PPFD) by covering or not covering the whole trees. Under a natural photoperiod (uncovered leaves), B-Put content and TGase activity paralleled the diurnal PPFD pattern, reaching a maximum at the highest PPFD; prior to this maximum, free PAs showed a significant rise. Plants that were in darkness until midday and suddenly exposed to high light intensity showed enhanced TGase activity, resulting in the maximum accumulation of B-Put. The involvement of the accumulation of B-Put reflected in the changes of the B-Put/bound spermidine ratio during the photoprotective responses to high light stress in forest plants is discussed in relation to the chlorophyll fluorescence parameters observed.

Water relations of seedlings of three Quercus species: variations across and within species grown in contrasting light and water regimes.

We compared seedling water relations of three Mediterranean Quercus species (the evergreen shrub Q. coccifera L., the evergreen tree Q. ilex L. subsp. ballota (Desf.) Samp. and the deciduous or marcescent tree Q. faginea L.). We also explored seedling potential for acclimation to contrasting growing conditions. In March, 1-year-old seedlings of the three species were planted in pots and grown outdoors in a factorial combination of two irrigation regimes (daily (HW) and alternate day watering (LW)) and two irradiances (43 and 100% of full sunlight). At the end of July, predawn and midday water potentials (Psi(pd), Psi(md)) were measured, and pressure-volume (P-V) curves were obtained for mature current-year shoots. Species exhibited similar Psi(pd) and Psi(md) values, but differed in leaf morphology and water relations. The evergreens possessed larger leaf mass per area (LMA) and were able to maintain positive turgor pressure at lower water potentials than the deciduous species because of their lower osmotic potential at full turgor. However, the three species had similar relative water contents at the turgor loss point because Q. faginea compensated for its higher osmotic potential with greater cell wall elasticity. Values of Psi(pd) had a mean of -1.12 MPa in LW and -0.63 MPa in HW, and Psi(md) had a mean of -1.13 MPa in full sunlight and -1.64 MPa in shade, where seedlings exhibited lower LMA. However, the P-V curve traits were unaffected by the treatments. Our results suggest that Q. faginea seedlings combine the water-use characteristics of mesic deciduous oak and the drought-tolerance of xeric evergreen oak. The ability of Q. coccifera to colonize drier sites than Q. ilex was not a result of higher drought tolerance, but rather may be associated with other dehydration postponement mechanisms including drought-induced leaf shedding. The lack of treatment effects may reflect a relatively low contrast between treatment regimes, or a low inherent responsiveness of these traits in the study species, or both.

Association between radionuclides (210Po and 210Pb) and antioxidant enzymes in oak (Quercus coccifera) and mastic tree (Pistacia lentiscus).

The activity levels of naturally occurring radionuclides Polonium-210 and lead-210 in different subjects including plant species have direct or indirect impact on human beings. High levels of ionising radiation cause oxidative stress and the interaction between antioxidative defense and radionuclides is not well established in plant systems. In this study, we aimed to understand the impact of oxidative stress caused by 210Po and 210Pb in two Mediterranean plants; Quercus coccifera and Pistacia lentiscus. We analysed the constitutive and seasonal levels of 210Po, 210Pb, lipid peroxidation levels, superoxide dismutase (SOD) and ascorbate peroxidase (APX) activities in the field-collected samples. The highest activity concentrations of 210Po and 210Pb were detected in both plants in summer and Q. coccifera had higher levels than that of P. lentiscus. SOD and APX activity trends were different between oak and mastic; as compared to P. lentiscus, Q. coccifera efficiently used the two major components of antioxidative defense. Lipid peroxidation levels were low in both plants in all seasons except that of spring which were in good agreement with high antioxidant enzyme activities. In conclusion, we found that high 210Po and 210Pb activity concentrations in oak and mastic did not interfere with their growth and life cycles. The ability of both plants for survival and adaptation to Mediterranean environmental constraints provided an additional advantage for coping radionuclide induced oxidative stress as well.